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SAN FRANCISCO--The Green Bay Packers sure did a good job of figuring out the San Francisco 49ers' read-option attack.

After spending a considerable amount of time working on it in the offseason, they completely shut down that aspect of the 49ers' offense Sunday in the regular-season opener at Candlestick Park.

They did not, however, have an answer for wide receiver Anquan Boldin.

In a 34-28 loss, the Packers held 49ers quarterback Colin Kaepernick to 169 fewer yards rushing then he gained in their divisional playoff loss eight months ago. And in total, they held the 49ers to 90 yards rushing on 34 carries, a 2.6-yard average.

What they couldn't do was construct a plan that both stopped the read-option and contained Kaepernick and his modest group of offensive targets. The Packers poured so much into keeping Kaepernick from doing what he did to them last time, they forgot about the part where he can complete 27 of 39 passes for 412 yards and three touchdowns.

“We kept the quarterback in front of us and we stopped the run,” cornerback Tramon Williams said. “First thing you always want to do is stop the run. That's base. Obviously, it opened up some other things.

“They're a good team. We knew they were going to make some plays. We were just trying to limit those plays.”

It didn't happen nearly enough.

The Packers secondary took a hit when safety Morgan Burnett (hamstring) couldn't make it through the week of practice and wound up being inactive. It forced defensive coordinator Dom Capers to go with Jerron McMillian and M.D. Jennings, two nominal starters, to patrol the back end of the secondary.

It turned out to be far worse than the Packers could have imagined.

Whereas Burnett has mostly overcome his propensity for mental mistakes and become aggressive to the ball, both McMillian and Jennings appeared tentative and missed out on a couple of big hits that might have sent a message to 49ers receivers. They allowed the 6-foot-1, 220-pound Boldin to outfight them for ball after ball.

“I think when our safeties watch the film they are going to wish they made more plays on the ball,” McCarthy said. “There were a few times when the ball was pushed vertically and was in the air for enough time for us to possibly make that play.

“This is not the way we intended to come here today. We were confident in our play. We let them have too many things.”

Boldin finished the game with 13 catches for 208 yards and a touchdown. He was five yards shy of moving into the No. 2 position in the Packers' record book for opponent single-game receiving yards and nine yards shy of his all-time high of 217 against Detroit in 2003.

The Packers played a variety of coverages during the game, but most often they appeared to be in a zone that allowed all the defensive backs to be facing Kaepernick. Capers did not want the 49ers quarterback breaking into the free like he did in January when the Packers played a lot of man-to-man.

The problem was that the 32-year-old Boldin eats up zone coverage. He's not fast and so the cornerbacks didn't have to respect his deep routes much, but when the ball was in the air across the middle no one beat him to it Sunday.

“He did just a couple routes the whole time,” cornerback Sam Shields said. “There wasn't anything different. He was just more aggressive to the ball and catching the ball more.”

And the Packers didn't put any big hits on him nor did they provide the kind of pass rush they needed to help keep Boldin away from the ball. It appeared that Capers' plan required the cornerbacks to make sure Kaepernick didn't escape to the outside on read-option plays.

Whether that had an effect on how they played the receivers or whether they just didn't play zone well is something Capers is going to have to find out.

“Give them the credit,” cornerback Tramon Williams said. “The outside got focused so much on stopping the zone read...the guy's not just a running quarterback, he can throw the ball, too. He extends plays; guys are going to get open.

“According to our game plan, I thought we did a good job. We had a chance at the end. At the end of the day, that's what matters. A play here or there, we win the game.”

One of those plays was a pass that went through Williams' hands early in the game. If he catches it, the Packers have the ball deep in San Francisco territory at the very least.

But other than Shields knocking away a slant to Boldin, there weren't many contested passes. Shields had a couple of nice pass breakups, but he also missed a tackle on Boldin in the middle of the field that allowed him to go 25 more yards for a gain of 43 that set up a touchdown.

The play came less than a minute after running back Eddie Lacy had given the Packers a 28-24 lead midway through the fourth quarter. All of the momentum the Packers built was tossed away after Boldin made that play.

Williams has covered big receivers exclusively in previous games, but he said there was no option for that to be done in this game because of the scheme the Packers were playing. In other words, they were playing so much zone that there wouldn't have been all that many opportunities to follow him around anyway.

Boldin said 49ers coach Jim Harbaugh did a great job of dissecting what the Packers were doing on defense.

“”I think it was just watching them over there in the offseason,” Boldin said. “There were some plays in there that we schemed up just for them, that we thought would be there. And they were.”

Harbaugh said that his offensive line deserved a lot of credit for the big passing day because of the way it protected Kaepernick. The Packers did not blitz much and the front four of the Packers wasn't effective getting to the 49ers quarterback.

“I thought that the pass rush of the Packers was completely negated by our offensive line and our backs with the way they blocked and protected,” Harbaugh said.

It's hard to say if having Burnett or cornerback Casey Hayward (hamstring) in the lineup would have made a difference, but the 49ers weren't at full strength either. One can only imagine how good they'll be when star receiver Michael Crabtree returns sometime this year from a torn Achilles'.

If the Packers meet the 49ers again, they're going to have to do more than just stop the read-option.